Peter Frost's anthropology blog, with special reference to sexual selection and the evolution of skin, hair, and eye pigmentation

Thursday, May 14, 2009

The First European?

What did the first modern humans in Europe look like? The question comes up in a BBC2 series The Incredible Human Journey, which shows the reconstructed head of a man who lived in the Carpathian Mountains some 35,000 years ago. With its brown skin and broad nose, this ‘First European’ looks, well, very un-European.

The online article is followed by bitter comments. One observes: “It seems to me that this result was not modelled on actual human remains. But rather on the image the Ministry of Truth has on what a modern European OUGHT to look like.”

And my comment? First of all, Europeans look European because they have physical traits that are rare or absent in other human populations. Since these traits are specific to Europe, they probably developed there. And they would have done so only after the arrival of modern humans 40,000-35,000 years ago. Therefore, a European living 35,000 years ago should have looked a lot less European than the ones around today.

So much for the theory. What about the facts? I should first point out that the reconstructed head is based not only on the Carpathian cranium but also on other remains from the same period and even on non-skeletal data. Obviously, skeletal remains don’t preserve skin color. We know that early modern Europeans were darker-skinned because the alleles for white skin arose much later in time—about 11,000 years ago at the SLC45A2 (AIM1) gene and 12,000–3,000 years ago at the SLC24A5 gene (Norton & Hammer, 2007; Soejima et al., 2005). As a Science journalist observed: “the implication is that our European ancestors were brown-skinned for tens of thousands of years” (Gibbons, 2007).

Does this seem counter-intuitive? How could Europeans have been brown for so long and so far north? Isn’t white skin an adaptation to northern latitudes and low levels of UV light? Well, human skin is brown among indigenous northern Asians and Amerindians who live just as far north with the same UV at ground level. As I’ve argued elsewhere, the extreme depigmentation of Europeans is probably due to sexual selection, as evidenced by other unique color traits, i.e., diversification of eye and hair color (Frost, 2006; Frost, 2008).

Early modern Europeans probably had broad noses too. We see this not only in the Carpathian cranium but also in other cranial remains from the same time period. We see this especially in a pair of skeletons from Grimaldi, northern Italy. The skeletons were initially dated to the early occupation of Europe by modern humans, c. 30,000 BP. Associated artifacts have since been radiocarbon dated to 14,000-19,000 BP but may come from later occupation layers (Bisson et al., 1996).

The Grimaldi skulls don’t look European. The face is wide but not high, the nose broad and flat, the upper jaw forward-projecting, and the chin weakly developed. The well-preserved dentition is not at all European. Among currently living populations, the ones who most closely resemble the Grimaldi humans seem to be the Khoisan peoples of southern Africa. The French physical anthropologists Boule and Vallois (1957, pp. 290-291) describe these early Europeans as having an almost African phenotype:

Comparisons which we have been able to make with the material at our disposal, in particular with the skeleton of the Hottentot Venus [a Khoisan individual], have led us to note, for instance, the same dolichocephalic character, the same prognathism, the same flattening of the nose, the same development of the breadth of the face, the same form of jaw, and the same great size of teeth. The only differences are to be found in the stature and perhaps in the height of the skull.

We know less about the soft-tissue characteristics. Alongside the skeletons were a number of female statuettes with big breasts, protruding bellies, full hips, and large buttocks. On the statuettes, the hair seems to be short and matted (Boule & Vallois, 1957, p. 311).

When did this original phenotype disappear? The data increasingly suggest that Europeans assumed their present-day appearance relatively late and over a relatively short time span. This transformation essentially took place during the last ice age (25,000 – 10,000 BP), with most of the changes probably occurring during the period after the glacial maximum (15,000 – 10,000 BP). As I’ve argued elsewhere, the cause was probably an intensification of sexual selection of women, i.e., too many women had to compete for too few men. On the one hand, male mortality increased in relation to female mortality because men had to cover much longer hunting distances. On the other, polygyny decreased because women depended much more on men for food provisioning (Frost, 2006; Frost, 2008).

This transformation took place on a vast expanse of steppe-tundra—today the plains of northern and eastern Europe—where highly mobile herds of reindeer and other herbivores were almost the sole food source for humans. The new phenotype must have then spread outward, via gene flow. Interestingly, the old phenotype may have persisted in some peripheral populations, perhaps into late prehistory and even after. As Boule and Vallois (1957, pp. pp. 291-292) note:

‘In Brittany, as well as in Switzerland and in the north of Italy, there lived in the Polished Stone period, in the Bronze Age and during the early Iron Age, a certain number of individuals who differed in certain characters from their contemporaries’, in particular in the dolichocephalic character of their skull, in possessing a prognathism that was sometimes extreme, and a large grooved nose. This is a matter of partial atavism which in certain cases, as in the Neolithic Breton skull from Conguel, may attain to complete atavism. Two Neolithic individuals from Chamblandes in Switzerland are Negroid not only as regards their skulls but also in the proportions of their limbs. Several Ligurian and Lombard tombs of the Metal Ages have also yielded evidences of a Negroid element.

Since the publication of Verneau’s memoir, discoveries of other Negroid skeletons in Neolithic levels in Illyria and the Balkans have been announced. The prehistoric statues, dating from the Copper Age, from Sultan Selo in Bulgaria are also thought to portray Negroids. In 1928 René Bailly found in one of the caverns of Moniat, near Dinant in Belgium, a human skeleton of whose age it is difficult to be certain, but which seems definitely prehistoric. It is remarkable for its Negroid characters, which give it a resemblance to the skeletons from both Grimaldi and Asselar.

It is not only in prehistoric times that the Grimaldi race seems to have made its influence felt. Verneau has been able to see, now in modern skulls and now in living subjects, in the Italian areas of Piedmont, Lombardy, Emilia, Tuscany, and the Rhone Valley, numerous characters of the old fossil race.

A similar observation is made by Fleure (1945):

In a few places in Sweden, Britain, and France people have been noticed who show characteristics of the skull and face that remind one of late-Paleolithic man: these people are usually darker, in hair and eyes, than their neighbors; sometimes they even have swarthy skins. Although this fact may not have great weight in argument, it does hint that there has been depigmentation in this region. The many stories of golden hair and blue eyes suggest that sexual selection may have helped the change.

An ancient Norse poem, the Rigsthula, describes how the god Rig created a class of thralls who were black-haired, swarthy, and flat-nosed (Jonassen, 1951). This theme comes up elsewhere in Old Norse literature (Karras, 1988). Thus, even in northern Europe, and as late as the proto-historic period, some Europeans may have retained a dark-skinned and broad-nosed phenotype. The ‘First European’ seems to have stayed around for a long time …

Norton, H.L. & Hammer, M.F. (2007). Sequence variation in the pigmentation candidate gene SLC24A5 and evidence for independent evolution of light skin in European and East Asian populations. Program of the 77th Annual Meeting of the American Association of Physical Anthropologists, p. 179.

It's consistent with the other 'Venus' figurines from the Upper Paleolithic (except that the head is absent).

It's misleading to ask which living race early modern Europeans belonged to. A genome study would probably associate them most closely with present-day Europeans. But overall changes to the genome are, at best, loosely correlated with changes to morphology (which involve only a fraction of the genome).

Morphologically, early modern Europeans seem to have been most similar to Khoisans, while being larger in body size and perhaps more adipose (although large fatty buttocks are also characteristic of Khoisan women).

Tod,

What strikes me about Venus figurines is that they all seem to have matted 'peppercorn' hair. The classic example is the Venus of Willendorf. The Venus of Brassempouy has longer hair than the others although her hair form still seems clumpy and not straight. She's later in time than the others (26,000-22,000 BP) and may correspond to the initial phenotypic changes of the last ice age (e.g., lengthening of head hair and narrowing of the nose).

"It's consistent with the other 'Venus' figurines from the Upper Paleolithic (except that the head is absent)."

Right. The Venus figurines all look like offensive linemen with upward sloping breasts.

"more adipose"

Does it ever occur to you to use Google?

The ancestors of modern Europeans arrived in Europe at least 40,000 years before present. Pre-glacial maximum Upper Palaeolithic males (before 16,000 BC) were tall and slim (mean height 179cm, estimated average body weight 67 kg), while the females were comparably small and robust(mean height 158 cm, estimated average body weight 54 kg).Stature of early EuropeansBased on these estimates, the mean BMI of UP European females was 21.6 (which is obviously not compatible with Venus figurines being depictions of typical females).

They are wearing hats. You can see the straight hair sticking out from beneath one of them.http://mathildasanthropologyblog.files.wordpress.com/2008/04/venus-hat.jpgI'd say the Venus of Brassempouy was wearing a woven hood, you can see in the close up near the bottom where the hair is beneath the hood.http://upload.wikimedia.org/wikipedia/commons/2/26/Venus_de_Brassempouy.jpgAnd they are definitely wearing some kind of plaited gear- most aren't butt naked.

A lot of those figurines just look fat rather than showing a San-type behind. The one in the link is an example. The male ivory head also has distinctly straight hair.

Caucasian type hair was known in the far East (Ainu have it)- this would suggest it predates the split of Asian and non Asian populations. You see straight hair on the carved male head(~26k bp).http://mathildasanthropologyblog.files.wordpress.com/2008/09/headbrugar11.jpg

As for the original posts- some Meso/Neolithic crania show some African traits (last time I checked the texts about 1/7) as they were contributed to partially by the Natufians who can trace their ancestry back to Nubia about 24k BP. They mixed in with the Turkish population and you see them in some Greek and Balkan sites. It's probably how the M78 Y chromosome wound up in SE Europe. These people had left Africa about 10k prior and were a minority in the Mesolithic and Neolithic expansions from Turkey.

My main issue with the reconstruction is that it is TOO dark, not that it IS dark; a lighter Khoisan yellow-brown would have made more sense- and isn't outside the range of people Arabia to Pakistan. These people didn't walk straight from Africa into Europe, a significant amount of time passed between the two events, and they'd have no reason to maintain so equatorial a skin colour. I am also familiar with the estimated dates for brown/fair and auburn hair, and these predate that crania quite significantly.

I'd question the Grimaldi looking like Khoisan. The last detailed description I read said that they had some typically European cranial traits (Keith).

Another source.. " The Grimaldi child was no more Negroid than the Palestinians of Skhul and many living Europeans of the Mediterranean region."(Coon, 1962)

The Natufians got described as 'negroid' too, but Loring Brace's study of them showed them to be essentially an mix of Eurasian and African, so any description of a crania as Negroid in older texts need to be taken with a pinch of salt, as they are often not that accurate and are aplied to any skull with a wider than average nasal aperture and any noticable prognathism.

As for the average early European..

"The results presented in Table 1 are consistent with the idea that Upper Paleolithic crania are, for the most part, larger and more generalized versions of recent Europeans. Howells ([1995]) reached a similar conclusion with respect to European Mesolithic crania."

There's also the issue of how accurate the reconstruction was and how complete the crania was to start with, as this can really affect the outcame.

The Grimaldi crania appear to be outliers in the normal range of UP Europeans. I suspect this crania also was.

As a side note about the figurines. They must have actually seen an obese woman at some point to have represented them so well. Maybe they specially fattened up the odd female for ritual purposes.

Didn't mean to sound dismissive Mathilda. I agree the Venus of Willendorf looks more like a hat than "matted 'peppercorn' hair", but I think it looks most like certain styles of cornrows.

The Venus of Willendorf could be a a heavy ribbed fabric hat explaining the cornrow effect. The direction of the ribbed pattern isn't conclusive either way. Ribbed fabric would be rather advanced for this period though.

To me the horizontal incisions in the hair/hood of The Venus of Brassempouy look like an attempt to depict a primitive tartan-like grid pattern in a fabric. Simple vertical lines would make it hair, but with the grid pattern I'm not sure.

The background on string skirts you give in your post is convincing so yes "they are definitely wearing some kind of plaited gear".

My difficulty is more with the context. It is clear what interest the figurines are intended to gratify. Usually the artist emphasized the primary sexual characteristic (generally deeply incised) and the secondary ones. Seen in this light the 'plaited gear', revealing as it is, may well have been a kind of lingerie.

For women covering the hair is more a sign of modesty as with conservative religions today, the sight of it is for the husband when behind closed doors. 'Let your hair down has a connotation. For these artists to render overtly sexual figurines while keeping their hats on seems like an anomaly.

The Venus of Brassempouy is different, i.e. not overtly sexual. It may depict a young maiden if so a hood makes sense.

I never seen such individuals with african atavisms but I've not doubt it's true.During his genetic research, Bryan Sykes tells the story of twins with a neanderthal-look in the UK, why not ? after all, if you can retrieve 40000 years old features, why not 200 000years old features ?

Could we also assume that these atavisms were more common 2000 years ago than today and were made of lots of legend?

Other than that, I am convinced by Peter Frost explanation of sexual selection, it makes perfect sense, but the real problem, to me, is not to explain how the modern european genes were selected, but how these genes spread trough thousands of miles in isolated tribes, even if we assume there was some sort of prehistoric trade. Because we know pre-modern europeans of the 9000-12000 years old range have many descendants today, all of these premodern europeans had to acquire the modern genes from other tribes since these genes are younger. I admit I don't get it.

"The BMI given is calculated while assuming they were not obese I think."

The BMI was calculated by me based on stature and body mass estimates given, which presumably are based on skeletal remains. It's possible to estimate body mass based on either skeletal size or stress. Results from these different methods tend to agree.

If it wasn't clear, this is merely the first paper I found containing these sorts of estimates. Other sources should be consulted, but if there were any evidence morbid obesity was common among Upper Paleolithic Europeans, I'm sure I would have heard about it. The point is: Frost should take a few minutes and check his speculation against the skeletal evidence.

It's impossible to estimate BMI simply from skeletal remains. You can have varying thicknesses of subcutaneous tissue with the same skeletal frame.

Mathilda,

1. I agree that head hair lengthened and straightened before the Caucasoid/Mongoloid split. This probably happened during the early portion of the last ice age (25,000 - 20,000 BP) when a common Eurasian population inhabited the steppe-tundra belt from southwestern France to Siberia. The advent of non-black hair colors probably postdates this split, since it is specific to European populations.

2. The Venus figurines in the Boule & Vallois textbook (p. 329 in the French edition) have large buttocks, often extremely large ones.

3. There is a lot of confusion with terms like 'African' and 'Negroid'. The earliest modern humans were probably Khoisan-like in appearance, but they didn't have many of the physical traits we now associate with the term 'Negroid' (black or dark brown skin, robust body build and muscular development, high incidence of polygyny, etc.). 'Negroid' populations seem to have originated in West Africa as a later evolutionary development, perhaps in association with the advent of female-dominated agriculture about 10,000 years ago in the Upper Niger valley.

4. I agree that the reconstruction is probably too dark.

5. I could quote many authors, like Coon, who dismiss the African-like appearance of the Grimaldi skeletons, often in a line or two. I think Boule & Vallois showed conclusively that this African appearance is not due to faulty reconstruction (which in any case could not explain the African-like dentition).

6. I don't think the Grimaldi skeletons are outliers. In a study of prehistoric European skeletons, Holliday (1997) found that early Upper Paleolithic skeletons had ‘tropical’ body proportions and clustered with recent Africans. Late Upper Paleolithic and Mesolithic skeletons clustered with recent Europeans. The shift to a more European phenotype is placed by Holliday (1997) at around 20,000 BP— during the last ice age and well after the arrival of modern humans.

7. "I am also familiar with the estimated dates for brown/fair and auburn hair, and these predate that crania quite significantly." Can you give a source? I follow the literature on the genetics of hair color and I've yet to see any reliable estimates for the origin of non-black hair colors. Harding has tried at different times to come up with a ballpark estimate, but she's now mum on the topic.

Tod,

To be honest, I can't really say for sure. It could be some kind of headdress. On the other hand, the pictures in Boule & Vallois look like hair (and why would the rest of their bodies be butt naked?)

RG,

I don't believe that this phenotypic transformation originated within all of Europe, just on the steppe-tundra plains of the north and east. During the last ice age, these plains were inhabited by migratory bands, so any new alleles for hair and eye color could have quickly spread within this zone.

The new phenotype then spread outward over a much longer period. I suspect that even within proto-historic times there were still some groups here and there with unusually dark skin and a more African facial morphology.

At the time of its relase in 1981, this movie was trashed by most school teachers as a completely irrealistic fairy tale . Even me, as a kid, I found completely stupid the fact that the main heros, an european man of the ice age, was able to travel in a couple of days to what I though had to be Africa, since he met people of obvious african appearance.Needless to mention the neanderthals that were supposed to be dead 10000 years earlyer...I thought.Now, give it justice, this movie might have been the best in its genre, more than anybody had thought before.

Same about the albino slave with a Down syndrome appearance, or the blue eyes of the almighty God of the pyramid in "10000 BC". I wouldn't trash them, because maybe even indirectly, such individuals were also part of the whole story, for exemple by pushing the trends of sexual selection to european-like features, before these features actually appeared.

A prehistory of the north: human settlement of the higher latitudes (2005)By John F. Hoffecker [p101]

"Climates ameliorated rapidly after 16,000- 15,000 years ago and new changes began to sweep across western Europe. Sites reappeared in northern France, Belgium, northwest Germany and southern Britain between 15,500 and 14,000 years ago. Although many of them are considered late Magdalenian, other industries emerged in the northern areas containing distinctive curve-back and tanged stone points. An emphasis on reindeer hunting persisted through a cold oscillation during 13,000-11600 years ago (Younger Dryas event). Deglaciation resumed after this cold event..."

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Dr. Manning's book mentioned the Tibetans being light skinned for their UV exposure in connection with their practice of polyandry

"Comparisons between measured and calculated UV-B dose rates on clear-sky days in Lhasa show good agreement. Comparisons of UV-B radiation levels in Lhasa (Tibet), Oslo (Norway), and Dar-Es-Salaam (Tanzania) show that the UV-B dose rates during the summer in Lhasa are higher than the maximum value in Dar-Es-Salaam, which is at the sea level in the equatorial region, and 60% higher than in Oslo, which is at the sea level but 60 degrees North. We conclude that the UV-B dose rates during the summer on the Tibetan plateau are among the highest levels in habituated regions of the world. Maximum measured daily-integrated UV-B doses in Lhasa range from about 10 kJ/m2 in the winter to about 65 kJ/m2 in the summer."Ground-based measurements and modeling of solar UV-B radiation in Lhasa, Tibet

You seem to be making the following argument: 1) fatty tissue imposes a weight load on the skeletal frame; 2) natural selection will compensate by increasing the width and density of certain bones, like the femur; 3) therefore, it's possible to estimate the adiposity of an extinct human by examining the width and density of the supporting bones.

Frankly, I'm not convinced (nor am I convinced by appeals to authority). First, the skeletal frame has to support not only fatty tissues but also muscle (which is denser and heavier than fat). Any estimate of adiposity would therefore be dependent on assumptions about muscle thickness (which also varies significantly). Second, and more importantly, it is doubtful that variations in bone width would be able to capture variations in fatty tissue thickness, all the more so because we're dealing with small samples of skeletal remains whose age and sex are often conjectural.

Dr. Frost -- I am a layman. Your view, as I understand it, is that the ancestors of present-day Europeans were little different from present-day Khoisan 25,000 years ago -- maybe even more recently. In "The Origin of Races," Carleton Coon classed the Khoisan and Caucasians as separate races. He expressed the view that the differences between human races are too great to have evolved recently, and he suggested that they go back at least to Homo erectus. Maybe there are living anthropologists who hold such a view. (I don't know.) I will appreciate your commenting on this. -- Philadelphia, Pennsylvania, USA

PS It has been a long time since I read "The Origin of Races." I state the above from memory.

You're referring to the multiregional model of human evolution. I only know of a few anthropologists who still subscribe to it (C. Loring Brace, Milford H. Wolpoff). The arguments against the multiregional model are archaeological and genetic:

1. Archaelogical data show a dramatic shift in Europe from Neanderthals to modern humans some 35,000 years ago. It doesn't look like modern humans evolved out of Neanderthals.

2. Both nuclear DNA and mitochondrial data have been retrieved from Neanderthal remains. Again, it doesn't look like modern humans evolved out of Neanderthals (although low levels of intermixture between the two groups cannot be excluded).

There are nonetheless many people who support a "hybrid model" where most humans today are the descendants of a group that spread out of Africa some 50,000 years ago but which also intermixed with archaic humans, like the Neanderthals in Europe.

I discuss the relative merits of the multiregional model and the Out-of-Africa model in my last post.

Thank you for your reply. You’ve clarified a number of things. I’ve read your new post, about the comparative strengths of the Out-of-Africa and multiregional views; but I find that I personally keep returning, in my mind, to one subject: the present-day differences among the races. Let us put aside for a moment paleoanthropology’s various types of genetic evidence. Let us also not ask, for the moment, what was the predecessor of European Homo sapiens. (I think Coon acknowledged that there was no fossil evidence of a European Homo erectus -- a major problem for his theory -- and I don’t think he argued that present-day Europeans were descended from Neanderthals.) I would just like your reaction to the apparent physical differences between, say, Bushmen and present-day Europeans. Is it possible that those differences evolved in the comparatively-short time your entry, above, suggests they have? It is difficult for me to say because, in a sense, I have nothing with which to compare them. I don’t know how long a period the fossil record suggests has been required for comparable differentiation in non-human species. I simply return to statements made by persons who have professional experience in this field, as you do. Coon, as I say, remarked almost offhandedly that the differences among what he saw as five human races (Caucasian, East Asian, Negro, Khoisan, and Australian) are too great to have evolved recently. In reading some version of the now-old UN statement “The Race Question,” I got the clear impression that, at the middle of the twentieth century, there were anthropologists who were not sure the present-day human population should even be regarded as one species. I presume those scientists were comparing human differences to differences found in other animals. To me, the races seem very different. When I consider that several million years are supposed to have been involved in the emergence of present-day humans from their most-recent non-human ancestor, I am struck that Out-of-Africa and your entry above suggest that the racial differences throughout the world have arisen in maybe only tens of thousands of years. Thank you again, Dr. Frost. You will understand that I appreciate the opportunity to discuss these subjects with a professional.

Significant differences in appearance and behavior can evolve in as little as 10 generations, if natural selection is strong enough. Physical change = natural selection X time. It is not simply a function of time alone.

If we look back over the fossil record, we see many examples of organisms that have remained unchanged for millions of years. We see others that seem to have changed very rapidly. (This is, incidentally, why we often have trouble finding transitional forms).

As to why human populations are so diverse in physical appearance, one reason is that our species has spread into a wide range of environments over the past 50,000 years. Much of this diversity thus reflects adapations to differences in climate or ecological setting.

Other differences seem to have been shaped by sexual selection. In a recent paper, I argue that sexual selection has increased the physical diversity of our species by acting more strongly on some populations than on others.

The paper can be read online at:

http://ww.jsecjournal.com/NEEPSfrost.pdf

I also recommend a recent and very readable book: "The 10,000 Year Explosion: How Civilization Accelerated Human Evolution" by Gregory Cochran and Henry Harpending. In it the authors argue that much human evolution has occurred over the last 10,000 years because cultural innovations have created new environments of adaptation.

Thank you, Dr. Frost, for another very-helpful answer. As soon as I read it, I visited jsecjournal.com, where I read your informative and intriguing article. I will be following your recommendation that I read "The 10,000 Year Explosion."

People do spend too much time considering the appearance of long dead Europeans where no real proof exists. Europe contained a number of human types in the past: Neanderthals, Cro Magnon modern humans, Mladec modern humans and others including the Grimaldi modern humans. Putting flesh to their bones would produce a wide array of looks, phenotypes which would look totally unrelated. The Cro Magnons are often described as anatomically modern humans exactly like today's Europeans in skeletal structure, but, those humans had traits all their own, specifically an elongated frame with long limbs, unusual hand shapes, a dysharmonic head shape not found in modern Europeans, a long headed skull with an extremely short face. In short, the Cro Magnons did not look like modern Europeans.

Dark pigmentation is the norm for humans. Dark hair, dark eyes, and a brownish skin. Those traits are found in humans from the tropics to the poles in all human types. The fact that some Europeans are atypical or abnormal compared with what appears to be a combination of physical traits found in most humans which has proven to be of survival value, is not particularly interesting. No more so than blond Australian Aborigines. Green eyed people are sporadically found in all racial types, as is the occasional reddish hair person. The European excess of fairness is just one of those strange things that happen to populations which have suffered population reduction followed by genetic drift. Nothing special.

What is more interesting is that the European population went through a severe population reduction due to the Black Death in historic times. Two thirds of the population of Europe died. That one event in recent history probably had more effect on the physical looks of Europeans today than sexual selection or adaptation to climate. It would have changed the European population more effectively and in a much shorter time frame than 11 thousand years ago for the mutation of genes that modify the effects of melanin in Europeans.

Forget those ancient remains. We are their descendents but not like them.

Genetic data only supports the multi-regional evolution theory. Specifically with regards to the MC1R gene that influences skin and hair pigment the LCA between Europeans and Africans is about 1 MYA. This implies that the evolutionary split must have occured before this date, and possibly, taking into account other genetic data before 2 MYA.

"Both African and non-African data suggest that the time to the most recent common ancestor is ~1 million years and that the age of the global 314 variant is 650,000 years. On this time scale, ages for the Eurasian-distributed Val60Leu, Val92Met, and Arg163Gln variants are 250,000–100,000 years; the ages for African silent variants—Leu106Leu, Cys273Cys, and Phe300Phe—are 110,000–40,000 years. For the European red hair–associated Arg151Cys and Arg160Trp variants, we estimate an age of ~80,000 years;"

I don't consider MCR1 variants to be particularly relevant. There are plenty of red haired, and other colored monkeys around. Think of the Orangutans. I suppose they are long lost Irishmen and women! The fact is that pheomelanin is made before it is converted to eumelanin. The MCR1 variants are just errors in the conversion process, hence little eumelanin gets made. Red haired people are in effect freaks. I have seen plenty of Albinos and their white skin, white hair and pale blue eyes are very attractive. Beauty is in the eyes of the beholder.

This is my personal opinion: No living European is descended from any human or hominin that lived in Europe before the LGM. They are all extinct like the Neanderthals. The famous Old Man of Cro Magnon would look more like an Amerindian than any European, he probably had a similar skin tone. Modern Europeans are the descendants of immigrants from Asia, mostly the Middle East and also Central Asia, that came in waves after the LGM. These immigrants spread R1b, R1a, J, G, E and I and mtDNA H into Europe. A large immigration wave occurred in the Neolithic bringing farming and Middle Eastern plants and animals, and later in the Bronze age immigrants brought I.E languages and with a subgroup containing R1a1, blond hair/blue eyes and a pinkish skin tone.

Forget silly figurines made by the first humans in Europe, they are not the ancestors of any Europeans. So it matters not whether their skin was brown, olive, swarthy or black, and whether their noses were flat as pancakes.

Considering that r1a has now been proven to more or less originate in India, it is more likely that south asians and central asians (r1b) came to europe, it is not likely that those two haplogroups would have to do with the origin of lighter hair/eyes.

I remember reading in one of Fleure's articles that people resembling the Grimaldi phenotype could still be found in a part of Wales called the Plynlimon moorlands. He even showed an old Daguerreotype of a Welsh clergyman whose complexion was rather dark and whose features seemed quite atypical of Northwest Europeans. His face actually reminded me of some Australian Aborigines, particularly the southeastern type that Joseph Birdsell called "Murrayian".

The survival of this type in a few isolated communities makes sense in view of parallel cases in the Americas. The Botocudos of Brazil, the Pericues of Baja California, and the Fuegians are all thought to have retained elements of this pseudo-"Australoid" morphology right up until the time of European contact.

Can one still find people in Wales who resemble the ones Fleure described? Or have these last pockets of Paleolithic ancestry been washed out by population shifts within the last century? We can only wonder.

Peter, to my knowledge, all of the post-neanderthal Europeans that have been assigned to a race or equivalent 'cluster' have turned out Caucasian save for Markhina Gora and Grimaldi.

Although within European Upper Paleolithic Caucasians there already existed two clusters identified by Howells, who named them as lateral and linear, and which broadly match the Cromagnid/Aurignacid division of Eickstedt as well as Coon's division separating the 'Upper Paleolithics' from the Mediterraneans.

The supposedly 'Negroid' features of certain Europeans sound suspect though I'd be interested in knowing the Grimaldi dental traits. This is the kind of odontometry that is most valuable in determining race, and you failed to list any such evidence for the non-European nature of the Grimadi teeth. And other than the possibility of such evidence from a comparison of the jaw and teeth, there is nothing in the verbal description that you cited that I feel is specifically Khoesanoid or African.

The tropical limb proportions of Early Upper Paleolithic Europeans were under selective pressure just as were those of their Patagonian analogs inhabiting a similarly cold climate and hunting large mammals.

'Ponto' is wrong that the Cro-Magnons are so distinct from northern Europeans and people with a similar stature and 'disharmonious' skull shape are not uncommon in regions of Europe today. The Cro-Magnon type clusters with Berg, Austria despite its long skull and is close to Iberomaurusian people from the Epipaleolithic of North Africa.

The Plynlimmon type resembles Combe Capelle, which was then thought to be Paleolithic, rather than the Grimaldi type. It is not Australoid other than in its robustness, but a 'Coarse Mediterranean' similar to parallels in ie. Iberia and Sardinia.

Lastly if the description of the thralls created by Rig has an experiential basis, it is based upon peoples related to the Lapps.

The pre-Nordic inhabitants of Scandinavia are still recognised in the racial phenotypes of Scandinavians, for example rather obviously in the brachycephalic Strandids whose origins appear rooted in the Fosna Culture. The Lapps themselves are of course predominantly descended from Baltic natives of Mesolithic stock whose ancestors must have undergone a language shift to become Uralic speakers after contact with the Baltic Finns.

Needless to say that even the Lapps are Caucasian and not something akin to the Kalahari Bushmen.

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Welcome to my blog! For the most part, this page will be an extension of my website, with comments relating to my research. But it will also branch out into more general discussions of human evolution.